Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound...

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Beaked whales echolocate on prey

1. Digital tags were used to describe diving and vocal behaviour of sperm whales during 198 complete and partial foraging dives made by 37 individual sperm whales in the Atlantic Ocean, the Gulf of Mexico and the Ligurian Sea. 2. The maximum depth of dive averaged by individual differed across the three regions and was 985 m (SD = 124.3), 644 m (123.4) and 827 m (60.3), respectively. An average dive cycle consisted of a 45 min (6.3) dive with a 9 min (3.0) surface interval, with no significant differences among regions. On average, whales spent greater than 72% of their time in foraging dive cycles. 3. Whales produced regular clicks for 81% (4.1) of a dive and 64% (14.6) of the descent phase. The occurrence of buzz vocalizations (also called 'creaks') as an indicator of the foraging phase of a dive showed no difference in mean prey capture attempts per dive between regions [18 buzzes/dive (7.6)]. Sperm whales descended a mean of 392 m (144) from the start of regular clicking to the first buzz, which supports the hypothesis that regular clicks function as a long-range biosonar. 4. There were no significant differences in the duration of the foraging phase [28 min (6.0)] or percentage of the dive duration in the foraging phase [62% (7.3)] between the three regions, with an overall average proportion of time spent actively encountering prey during dive cycles of 0.53 (0.05). Whales maintained their time in the foraging phase by decreasing transit time for deeper foraging dives. 5. Similarity in foraging behaviour in the three regions and high diving efficiencies suggest that the success of sperm whales as mesopelagic predators is due in part to long-range echolocation of deep prey patches, efficient locomotion and a large aerobic capacity during diving.

Deep‐diving foraging behaviour of sperm whales (Physeter macrocephalus)

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.

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Sperm whale behaviour indicates the use of echolocation click buzzes 'creaks' in prey capture

Strandings of beaked whales of the genera Ziphius and Mesoplodon have been reported to occur in conjunction with naval sonar use. Detection of the sounds from these elusive whales could reduce the risk of exposure, but descriptions of their vocalizations are at best incomplete. This paper reports quantitative characteristics of clicks from deep-diving Cuvier’s beaked whales (Ziphius cavirostris) using a unique data set. Two whales in the Ligurian Sea were simultaneously tagged with sound and orientation recording tags, and the dive tracks were reconstructed allowing for derivation of the range and relative aspect between the clicking whales. At depth, the whales produced trains of regular echolocation clicks with mean interclick intervals of 0.43 s (±0.09) and 0.40 s (±0.07). The clicks are frequency modulated pulses with durations of ∼200 μs and center frequencies around 42 kHz, −10 dB bandwidths of 22 kHz, and Q3 dB of 4. The sound beam is narrow with an estimated directionality index of more than 25 dB...

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Echolocation clicks of free-ranging Cuvier's beaked whales (Ziphius cavirostris).

SUMMARY Blainville9s beaked whales ( Mesoplodon densirostris Blainville)
echolocate for prey during deep foraging dives. Here we use acoustic tags to
demonstrate that these whales, in contrast to other toothed whales studied,
produce two distinct types of click sounds during different phases in
biosonar-based foraging. Search clicks are emitted during foraging dives with
inter-click intervals typically between 0.2 and 0.4 s. They have the
distinctive form of an FM upsweep (modulation rate of about 110 kHz
ms -1 ) with a -10 dB bandwidth from 26 to 51 kHz and a pulse length
of 270 μs, somewhat similar to chirp signals in bats and Cuvier9s beaked
whales ( Ziphius cavirostris Cuvier), but quite different from clicks
of other toothed whales studied. In comparison, the buzz clicks, produced in
short bursts during the final stage of prey capture, are short (105 μs)
transients with no FM structure and a -10 dB bandwidth from 25 to 80 kHz or
higher. Buzz clicks have properties similar to clicks reported from large
delphinids and hold the potential for higher temporal resolution than the FM
clicks. It is suggested that the two click types are adapted to the separate
problems of target detection and classification versus capture of low
target strength prey in a cluttered acoustic environment.

Foraging Blainville's beaked whales (Mesoplodon densirostris) produce distinct click types matched to different phases of echolocation.

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vesteralen, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.

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Anders Lund

Papers

The monopulsed nature of sperm whale clicks